Journal of the Persian Gulf (Marine Science)/Vol. 4/No. 12/June 2013/8/23-30 Report on the Chromosomes of Liza klunzingeri (Day, 1888) in Zyarat Estuary, Persian Gulf Faqih Ahmadani, Ahmad1; Hosseini, Seyed Javad*2,3, Qasemi, Seyed Ahmad2; Mohammadi, Gholam Hossein4 1- Dept. of Aquaculture, Faculty of Agriculture and Natural Resources, Islamic Azad University, Ahvaz, IR Iran 2- Inistitute of Persian Gulf, Persian Gulf University, Busheher, IR Iran 3- Dept. of Cell and Molecular Biology, Faculty of Science, Persian Gulf University, Busheher, IR Iran 4- South of Iran Aquaculeare Research Center, Ahvaz, IR Iran Received: Febrmary 2012 Accepted: December 2012 © 2013 Journal of the Persian Gulf. All rights reserved. Abstract This paper reports on the karyotyping of mullet fish species, Liza klunzingeri, endemic of the Persian Gulf. The metaphase chromosome spreads, obtained from kidney cells of 13 specimens were examined. The diploid chromosome number was found to be 2n=48. The karyotype was discovered to be consisted of 48 acrocentric chromosome pairs. The arm number was determined to be NF=48. The chromosome number observed in this fish is the same as the majority of Mugil fish, thus confirming the commonality of this feature in the family. Keywords: Liza klunzingeri, Chromosome, Karyotype, Persian Gulf coasts of the Khuzestan province (Kashy et al., 2007). This fish is dispersed from the Persian Gulf to the Indian Ocean, Arabian Sea and the Gulf of Oman (Randall, 1995). Catching is done by beach seines, set nets, and gill nets (Ismail et al., 1998). Studies worldwide on Liza klunzingeri are done primarily based on morphological and anatomical traits, respectively (Ismail et al., 1998; Abou-Seedo and Dadzie., 2004; Kashi et al., 2007; Valinasab et al., 2005). Fish chromosome studies is one of the various applications of genetics in fishery research and has several applications in taxonomic studies, genetics, breeding and biotechnology. Many morphologically similar fish may have different types and numbers of 1. Introduction Liza klunzingeri belongs to Mugilidae family, which is distributed in various coastal aquatic habitats of the world’s tropical, subtropical and temperate regions. This family includes 17 genera and 72 species (Nelson, 2006). The fish habitat is various coastal substrata, brackish waters, and lagoons with high salinity (Golani et al., 2002). Because of the wide variety and small differences between mullet species, considerable divergences exist in their systematic classification. Liza klunzingeri is one of the mullet fish species that has valuable fisheries catch over the * Email: [email protected] 23 Faqih Ahmadani et al. / Report on the Chromosomes of Liza klunzingeri… identified (Table 1), yet there is no report on the type and number of chromosomes of Liza klunzingeri. The aim of this study is to determine the number and type of the chromosomes of Liza klunzingeri, along with its karyotype, the number of chromosomal arms (NF), bases along the chromosomes, and the ratios of long and short arms. chromosomes. Therefore, knowledge of normal karyotype of aquatic species can be instrumental in such studies. Before applying the method of breeding fish, acquiring enough information about the type and number of chromosomes of the species is necessary (Gold et al., 1990). Karyotype of many mullet fish species of the genus Mugil and Liza have been Table 1. Chromosome number in 19 Mugilidae species Species 2n Chromosomal formula NF Source Oedalechilus labeo 48 2St +46a 48 Rossi et al., 2000 Mugil cephalus 48 48a 48 Rossi et al., 1996 M.trichodon 48 48a 48 Nircho et al., 2005 M.speigleri 48 48a 48 Rishi and Singh., 1982 M.platanus 48 48a 48 Jordao et al., 1992 M.parsia 48 48a 48 Khuda-Buksh and manna., 1974 M.liza 48 48a 48 Nirchio and Cequea., 1998 M.gaimardianus 48 48a 48 Nirchio et al., 2003 M.curema 24 22m+2Sm 48 Nirchio and Cequea., 1998 M.corsula 48 48a 48 Khuda-Buksh and manna., 1974 M. curema 28 20m+4St+4a 48 Le Grande and Fitzsimons., 1976 Liza aurata 48 2St +46a 48 Choudry et al., 1979 L.saliens 48 2St +46a 48 Gornung et al., 2001 L.saliens 48 46a +2Sm 50 Arefyev., 1989 L.ramada 48 2Sm +46a 48 Rossi et al., 1997 L.ramada 48 46a +2Sm 50 Delgado et al., 1992 Paramugil parmatus 48 48a 48 Choudry et al., 1979 Chelon macrolepis 48 48a 48 Choudry et al., 1979 L.aurata 48 46a+2St 48 Cano et al., 1982 L.aurata 48 46a+2Sm 48 Delgado et al., 1991 L. saliens 48 46a +2St 48 Cataudella et al., 1974 L. ramada 48 46a+2St 48 Cataudella et al., 1973 L. ramada 48 46a +2St 48 Cataudella et al., 1974 L. ramada 48 46a +2Sm 48 Delgado et al., 1991 Chelon labrosus 48 2St+46a 48 Cataudella et al., 1974 Liza klunzingeri 48 48a 48 Present Study 48 46a+2st 48 Nirchio et al., 2008 48 48a 48 Hett et al., 2011 Agonostomus monticola M.incilis 24 Journal of the Persian n Gulf (Marin ne Science)/Vool. 4/No. 12/Ju une 2013/8/233-30 trreatments weere determinned to be 50 – 60 and 30 min, m respectiv vely. Both thhe types of sllide preparattion were w effectiv ve in obtainiing well-sprread metaphhase ch hromosomess. Analyses of chromo osome spreeads reevealed that modal chroomosome num mber containned 48 (2n=48) (T Table 2) (Figg. 1). 2. Materialls and Methods Thirteen specimens of Liza klunzingeri k were m Zyarat Estuuary and tran nsported alivee to a caught from well-aeratedd aquarium in the laborratory with room temperaturee, before chroomosomal an nalysis. Karyologgical study was w carried out accordinng to Nirchio aand Colleaggues (2005 5) with some modificationns. Each fishh was injectted with 25µ µgr/gr body weight ht colchicine solution, and d scarified aft fter 60 ± 10 min. K Kidney tissue was then rem moved and pplaced in physioloogical serum m, and cell suspension was obtained byy repeated pipetting. p Thee suspensionn was centrifuged at 1100 rpm for 10 min att 4 °C. Cell ppellets were treatedd by suspenssion in hypottonic 0.075 m molar KCl solutionn and tested for f 20, 25, 30 and 35 minuutes at room tempeerature. Sampples were then n centrifugedd, and hypotonic soolution was replaced r with h freshly preepared methanol-gllacial acetic acid a (3:1) as a fixative. Sam mples were fixed for 40 to 60 6 minutes with w 2 or 3 time changes eeach. The suspension obtained was centrifuged, and the supernatantt was rem moved. Afterwards, the cell pelleets were resusspended in thhe few remaining drops. For F chromosomal sppreads preparation, the cool celll suspension was droppedd onto pre-warmedd clean slide (45-50 ( °C). In another meethod, cell suspenssion was drropped onto cold clean slide. Staining waas performed with 10 and d 20% Gimsaa in a Sorenson buuffer (pH 6.8)) for 10 – 20 min. Microsscopic observationss were doone with a Nikon Light microscope. Total length th of each ch hromosome, short and long arm ms length annd arm ration n were determ mined using MicrooMeasure sooftware V3.3 according tto the Levan methood (Levan et al., 1964). Table 2. Frequencies distributiions of chromo osome counts from f kiidney cells of Liza L klunzingerii. Speciees Liza klunziingeri 45 3 46 4 2n 47 3 Totaal 48 64 49 2 76 3. Results Each of the steps innvolved in the chromoosome study is impportant in atttaining largee number off well spread metaaphase chrom mosomes. Am mong these ssteps, the colchiciine and hyppotonic treattment were more important. The optimaal colchicinees and hypootonic Fiig. 1: Chromoso omal Spreads (1100X) of Liza kllunzingeri, 2n = 48. 25 Faqih Ahmadani et al. / Report on the Chromosomes of Liza klunzingeri… data are summarized in Table 3. The chromosomal spreads obtained allowed us to propose that karyotype of Liza klunzingeri consist 48 acrocentric and NF=48 (Fig. 2). The ideogram of the L.klunzingeri was made on the basis of the karyotype (Fig. 3). The chromosomes of 76 chromosomal spreads were counted, and 64(84.2%) had 48 chromosomes. Chromosome number varied from 45 to 49 (Table 2). Figure 1a shows karyotype analysis using MicroMeasure V3.3. The morphological and numerical Table 3: Morphometric data of the chromosomes of Liza klunzingeri (2n=48). Relative lengths (RL) (% of the set) and centromeric indices (CI). No 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Length each 33.5086 32.2939 31.9234 30.0397 28.5103 28.3604 28.2633 27.5744 27.4587 27.3895 27.3293 27.3200 27.0466 26.9051 26.5476 26.4511 25.5193 25.3447 25.2866 25.2676 25.2364 24.4698 24.3814 24.0288 23.9604 23.8563 23.6541443 23.4824 23.4562 23.3939 23.3040 23.2080 23.0900 22.6092 22.2277 22.2101 22.1953 22.0883 21.8451 21.6987 21.1448 20.9747 20.8279 20.3817 19.7821 19.0369 17.8996 15.6710 Relative lengths % of set 2.8434% 2.7404% 2.7089% 2.5491% 2.4193% 2.4066% 2.3983% 2.3399% 2.3301% 2.3242% 2.3191% 2.3183% 2.2951% 2.2831% 2.2527% 2.2446% 2.1655% 2.1507% 2.1457% 2.1441% 2.1415% 2.0764% 2.0689% 2.0390% 2.0332% 2.0244% 2.0072% 1.9926% 1.9904% 1.9851% 1.9775% 1.9694% 1.9593% 1.9186% 1.8862% 1.8847% 1.8834% 1.8743% 1.8537% 1.8413% 1.7943% 1.7799% 1.7674% 1.7295% 1.6786% 1.6154% 1.5189% 1.3298% Long arm 33.5086 32.2939 31.9234 30.0397 28.5103 28.3604 28.2633 27.5744 27.4587 27.3895 27.3293 27.3200 27.0466 26.9051 26.5476 26.4511 25.5193 25.3447 25.2866 25.2676 25.2364 24.4698 24.3814 24.0288 23.9604 23.8563 23.6541 23.4824 23.4562 23.3939 23.3040 23.2080 23.0900 22.6092 22.2277 22.2101 22.1953 22.0883 21.8451 21.6987 21.1448 20.9747 20.8279 20.3817 19.7821 19.0369 17.8996 15.6710 26 Short arm 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Arm Ratio (L/S) N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A Cent. Index (S/(L+S)) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Classification Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Acrocentric Journal of the Persian n Gulf (Marin ne Science)/Vool. 4/No. 12/Ju une 2013/8/233-30 1 2 3 4 5 6 7 8 9 10 11 12 133 14 15 16 17 18 199 20 21 22 23 24 Fig. F 2: Karyotyppe (100X) Liza klunzingeri, 2n n = 48. Fig. F 3: Ideogram m of Liza klunzzingeri chromossomes h 30 min hyppotonic incub bation. Fixattive obtained with trreatment waas not founnd to be ass important as co olchicine an nd hypotoniic treatment in obtainning ch hromosome preparation;; but perso onal experieence sh howed that overnight o mai aintenance off cell suspenssion in n fixative so olution in ref efrigerator reesulted in beetter sllide preparatiion. The chrom mosome numbber of most of o the speciess in th he genus Liiza is 2n=448, 46 pairs of which are accrocentric. However, H alll 48 chrom mosomes of the Liza L klunzing geri in this sstudy, are accrocentric. In I a reeview study conducted bby Sola et al. a (2008), thhree Cytotypes C of chromosom mal formula are defined for mullet m fish: Cytotype C A, w which consists of 48 pairss of accrocentric ch hromosomes,, accounted for the greaatest number (10 species beelonging to four geneera); Cytotype C B which w consistts of 46 pairrs of acrocenntric ch hromosomes, plus twoo pairs of subtelocenntric 4. Discussioon Chromosome studies are basically similaar in methodologyy, almost all including colchhicine, hypotonic, ffixative treattments, slidee preparationn, and staining, rrespectively. A simplle method for Karyologicaal study of fish f is the diirect chromoosome preparation from kidneyy of live sp pecimens beecause kidney tissuue in most fishes contains the hematoppoietic elements, whhich provide numerous bllood cells in aactive division (Goold et al., 19990a). In this stuudy, colchicine treatmen nt was found to be important inn obtaining metaphase m ch hromosomess. The best time foor colchicinee treatment was w determinned to be 50 to 660 min. Decreasing tim me of colchiicines treatment reesulted in low l metaphaase chromossomes and longerr treatmentt led to more m condeensed chromosom mes. The besst chromosom me spreads were 27 Faqih Ahmadani et al. / Report on the Chromosomes of Liza klunzingeri… that great metacentric or submetacentric chromosomes have been generated through the reduction in number of the chromosomes by the Robertsonian fusion (Doucette and Fitzsimous., 1998). Diversity in the karyotype of the genus Mugil may be due to the centromere fusions of the acrocentric chromosomes (Le Grande and Fitzsimons., 1976). In a report by Legrande & Fitzsimons (1976), the number of the chromosomes of M.curema in the Caribbean was 2n=24, while in another report by Nirchio & Cequea (1998), this number for the same fish in the Gulf of Mexico was 2n=28. Since the arms of chromosomes in M.curema were longer than those in grey mullet and M.liza, it seemed reasonable that the karyotype of this fish might have evolved more than other species of the mullet family (Legrande and Fitzsimous., 1976; Nirchio and Cequea, ., 1998) Despite their systematic similarities, cytogenetic information distinguishes M. curema and M. rubrioclalus (Nirchio et al., 2007). Therefore, we can conclude that the mullet family, except for the M. curema, possess 48 pairs of chromosomes, most of which are acrocentric (Sola et al., 2007, 2008). Like all other species of the genus Liza, all the chromosomes of the Liza klunzingeri were determined to be acrocentric. Complementary information can be obtained through further research on this species in the Persian Gulf, Gulf of Oman, Indian Ocean. (observed in five species belonging to three genera) and Cytotype C which consists exclusively of biarmed chromosomes. M. curema from Louisiana and Brazil (2n=28, NF=48) and from Venezuela (2n=24, NF=48) display these cytotypes. The karyotype of Liza klunzingeri can be assigned to cytotype A. It is likely that other studies around the world on Liza klunzingeri and comparison of those studies yield different results. For example, the chromosomal formulae for L. Ramada was 2n= 48= 46a+ 2st (Cataudella et al.,1973, 1974); yet for the same species, Delgado et al. (1992) obtained 2n=48=46a+ 2sm. The same is true about L. saliens, for which the two chromosomal formulae were 2n= 48=46a+2st (Gornug et al., 2001; Cataudella et al., 1974) and 2n=48=46a + 2sm (Arefyev et al., 1989). Two different chromosomal formulae for L. aurata are reported, 2n=48= 46a+ 2sm (Cataudella et al., 1974; Delgado et al., 1992) and 2n=48=26a+ 2st (Cano et al., 1982). On the one hand, high concentration and longterm effects of colchicine causes high compression, shrinkage and shortening of the chromosome arms (Beck et al., 1980); This makes it difficult to detect the short arms of the chromosomes; and consequently, the mistakes in distinguishing different chromosomsl arms leads to differences observed in the karyotypes (Nirchio and Cequea., 1998). On the other hand, due to their small size and high compression, detection of chromosome arms is very difficult and sometimes it is impossible to detect homologous chromosomes. To overcome this problem, method of chromosome banding could be employed. Provided that the facilities are available, rating band method could be very useful in identifying the chromosomes more precisely; since without such patterns in band rating chromosomal arms, detecting the chromosomal similarities is only possible based on the chromosome size and the position of the centromere (Nirchio et al., 2008). Recent studies using chromosomal markers through realignment of chromosomes have shown that the Chromosomal evolution of mullet is more complex than it was believed (Sola et al., 2008). Karyologic study of twenty species of Clupeifromes and Elopifromes has shown Acknowledgements This work was granted by the Persian Gulf University, Grant of 19/63. We would like to thank all those who assisted in the field collection and laboratory processing of samples for this project. We are grateful to Mr. Ali Nakisa for editing the manuscript. References Abou-Seedo, F.S. and Dadzie, S., 2004. Reproductive cycle in the male and female grey mullet, Liza klunzingeri in the Kuwaiti waters of 28 Journal of the Persian Gulf (Marine Science)/Vol. 4/No. 12/June 2013/8/23-30 Mediterranean, In: Workshop on Lessepsian Migration Proceedings (Ozturk B. & N. Basusta, eds.). pp 1-9. Istanbul: Turkish Marine Research Foundation. Gold, J.R., Li, Y.C., Shipley, N.S. and Powers, P.K., 1990a, Improved methods for working with fish chromosomes with a review of metaphase chromosome banding. Journal of Fish Biology. 37:567-575 Gold, J.R., Jenkin, J.D. and Powers, P.K., 1990, Cytogenetic studies in North American minnows (Cyprinidae). XVIII Chromosomal NOR variation among eight species. Cytologia. 55:483-492. Gornung, E., Cordisco, C.A., Rossi, A.R., Innocentiis, S.De., Crosetti, D. and Sola, L., 2001, Chromosomal evolution in Mugilidae: karyotype characterization of Liza saliens and comparative localization of major and minor ribosomal genes in the six Mediterranean mullet. Marine Biology. 139: 55-60. Hett. A.K., Nirchio, M., Oliveira, C., Siccha, Z., Rossi, A.R. and Sola, L., 2011, Karyotype characterization of Mugil incilis Hancock, 1830 including adscription of an unusual co-localization of major and minor ribosomal gene in the family. Neotropical Ichthyology Neotropical Ichthyology. 9 (1): 107-112. Ismail, W.A.K., Al-Abdul-Elah. and Al-Yamani, F., (1998) Larval development of the back keeled mullet Liza carinata. Hydrobiologia 10-1998, 385 (1-3): 87-105 Jordão, L.C., Oliveira, C., Foresti, F. and Godinho, H., 1992, Caracterização citogenética da tainha, Mugil platanus (Pisces, Mugilidae). Boletim do Instituto de Pesca. 19:63-66.(In Portuguese) Kashi, M.T., Hashemi, A.R. and Safikhani, H., 2007, Study of some growth characteristics of Klunzingeri Mullet (Liza klunzingeri) in the coastal of Khuzastan, The full paper is in Persian and the abstract is available in English on http://www.sid.ir/fa/VEWSSID/J_pdf/3421387040 3.pdf the Persian Gulf. Cybium. 28(2):97-104. Arefyev, V.A., 1989, The application of the method of colchicines baths to studies of karyotype of the young of two mullet species (Mugilidae) from the black sea. In Dushkina LA. Early Life History of Marine Species. PP: 139-149. Sbornick Nauchin Trudov VNIRO, Moscow. Beck, M.L., Bingger, C.J. and Dupree, H.K., 1980, Caryological analysis of Ctenopharyngodon idella, Aristichthys nobilis and Their F1 Hybrid. Transactions of The American Fisheries Society.109: 133-438. Cano, J, Thode, G, and Alvarez, M C. 1982. Karyoevolutive considerations in 29 Mediterranean teleost fishes. Vie Milieu. 32: 21–24. Cataudella, S. and Capanna, E., 1973, Chromosome complements of three species of Mugilidae. Experientia. 29: 489–491. Cataudella, S., Civitelli, M.V. and Capanna, E., 1974, Chromosome complement of the Mediterranean Mullets. Carylogia. 27:93-105. Cervigón, F., 1993, Los peces marinos de Venezuela. Fundación Científica Los Roques. Vol. II. 2da Edición. Caracas, Venezuela. (In Spanish) Choudhury, R.C., Prasad, R.C. and Das, C., 1979, Chromosomes of six species of marine fishes. Caryologia. 32: 15–21. Delgado, J.V., Thode, G., Lobillo, J., Camacho, M.E., Alonso, A. and Rodero, A., 1991, Detection de la region del organizador nucleolar en chromosomas de la familia Mugilidae (perciformes): Precisones tecnicas, Archivos de zootecnia. 40:301-305.( In Spanish) Delgado, J.V., Molina, A., Lobillo, J., Alonso, A. and Camacho, M.E., 1992, Morphometrical study on the chromosome of three species of Mullet (Teleostei،Mugilidae). Carylogia. 45:263-271. Doucette, A.J.J.R. and Fitzsimons, J.M., 1988, Karyology of Elopiform and Clupeiform fishes. Copeia. 1: 124-130 Golani, D., 2002, Lessepsian fish migration: characterization and impact on the eastern 29 Faqih Ahmadani et al. / Report on the Chromosomes of Liza klunzingeri… of Hawaii Press, Honolulu, Hawaii. 439P. Rishi, K.K. and Singh., 1982, Karyology studies on five estuarine fishes. Nucleus 25:178-180. Rossi, A.R., Capula, M., Crosetti, D., Sola, L. and Campton, D.E., 1995, Allozyme variation and genetic differentiation among global populations of striped mullet Mugil cephalus (Pisces: mugilidae). Copeia (in Press). Rossi, A.R., Crosetti, D., Gornung, E. and Sola, L., 1996, Cytogenetic analysis of global Population of Mugil cephalus by Different Staining techniques and Fluorescent in situ hybridization, Heredity 76 (6):77-82. Rossi, A.R., Gornung, E. and Crosetti, D., 1997, Cytogenetic analysis of Liza ramada by Different Staining techniques and Fluorescent in situ hybridization. Heredity. 79:83-87. Rossi, A.R., Gornung, E., Crosetti, D.S., Innocentiis, De. and Sola, L., 2000, Cytogenetic analysis of Oedalechilus labeo (Pisces: Mugilidae), with a report of NOR variability. Marine Biology. 136:159-162. Sola, L., Gornung, E., Mannarelli, M.E. and Rossi, A.R., 2007, Chromosomal evolution in Mugilidae, Mugilomorpha: an overview, Pp. 165-194. In: Pisano, E., Ozouf-Costaz, C., Foresti, F. and Kapoor, B.G. (Eds.), Fish Cytogenetic. Enfield, Science Publishers 502p. Sola, L., Nirchio, M. and Rossi, A.R.,2008, Past and future of cytogenetic of mugilidae: an updated overview. Boletin Del Instituto Oceanogr fico de Venezuela. 47: 25-31. Valinasab, T., Seyfabadi, S.J., Javadzadeh, N. and Safikhani, H., 2005, Reproduction of Klunzingeri mullet, Liza klunzingeri, in coastal waters of Khuzestan Province (Persian Gulf). Iranian Journal of Fisheries Sciences. 6(2);129-142. Khoda-Bukhsh, V.S. and Manna, G.K .,1974, Somatic chromosomes in seven species of teleostean fishes. Chromosome Information Service. 17: 5-6. LeGrande, W.H. and Fitzsimons, J.M., 1976, Karyology often mullets Mugil curema and Mugil cephalus (Perciformes: Mugilidae) from Louisiana, Copeia, 1976(2):388-391. Levan, A., Fredga, A. and Sandberg, A., 1964, Nomenclature for centromere position on chromosomes. Hereditas. 52: 201-220. Nelson, J.S., 2006, Fishes of the World, fourth ed. John Wiley and Sons, Inc, New York, NY. 624 P. Nirchio, M. and Hernan Cequea, T.Y., 1998, Karyology of Mugil Liza and M. Curema from Venezuela, Boletín de Investigaciones Marinas y Costeras.27:45-50. Nirchio, M., Gon, F.C., Porto, J.I.R., Perez, J.E., Gomez, J.A. and Vilalaz, J.,2003, Karyotype Supporting Mugil Curema Valenciennes, 1836 and Mugil gaimardianus Desmarest 1831 as two wiled nominal species. Scientia Marina. 67(1):113-115. Nirchio, M., Ron, E. and Rossi, A.R., 2005, Karyology Characterization of Mugil Trichodon Poey, 1876 (Pisces: Mugilidae). Scientia Marina. 69(4):525-530. Nirchio, M., Oliveira, C., Ferreira, I.A., Perez, J.E., Gaviria, J.I., Harrison, I., Rossi, A.R. and Sola, L., 2007, Comparative cytogenetic and allozyme analysis of Mugil rubrioculus and M. Curema from Venezuela. Interciencia. 32:757-762. Nirchio, M., oliveira, C., Irani, A., Martin, F.C., Rossi, A.R and Sola, L., 2008, Classical and molecular cytogenetic characterization of Agonostomus monticola a primitive species of Mugilidae (Mugiliformes). Genetica DOI 10. 1007/s10709-008-9256-8. Randall, J.E., 1995, Costal Fishes of Oman, University Faqih Ahmadani et al. / Report on the Chromosomes of Liza klunzingeri… Journal of the Persian Gulf (Marine Science)/Vol. 4/No. 12/June 2013/8/23-30 Journal of the Persian Gulf (Marine Science)/Vol. 4/No. 12/June 2013/8/23-30 30
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